New taxonomic and faunistic data on the funnel-weavers (Araneae, Agelenidae) of Turkiye and the Caucasus, with five new species

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ZooKeys 1218: 251-286 (2024)
DOI: 10.3897/zookeys.1218.135249

Research Article

New taxonomic and faunistic data on the funnel-weavers (Araneae,
Agelenidae) of Turkiye and the Caucasus, with five new species

Alireza Zamani'™®, Rahsen S. Kaya2®, Yuri M. Marusik?45©

on fF wo NY —

Zoological Museum, Biodiversity Unit, Fl-20014 University of Turku, Turku 20500, Finland

Department of Biology, Faculty of Arts and Science, Bursa Uludag University, TR-16059, Bursa, Turkiye
Department of Zoology & Entomology, University of the Free State, Bloemfontein 9300, South Africa
Altai State University, Lenina Pr., 61, Barnaul, RF-656049, Russia

Institute for Biological Problems of the North, Portovaya Str. 18, Magadan 685000, Russia

Corresponding author: Alireza Zamani (zamani.alireza5@gmail.com)

OPEN Qaccess

Academic editor: Dragomir Dimitrov
Received: 21 August 2024

Accepted: 27 October 2024
Published: 21 November 2024

ZooBank: https://zoobank.
org/7F6A7B71-74A9-42BA-A258-
C28544EAC887

Citation: Zamani A, Kaya RS, Marusik
YuM (2024) New taxonomic and
faunistic data on the funnel-weavers
(Araneae, Agelenidae) of Turkiye and
the Caucasus, with five new species.
ZooKeys 1218: 251-286. https://doi.
org/10.3897/zookeys.1218.135249

Copyright: © Alireza Zamani et al.

This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).

Abstract

New taxonomic and faunistic data on the agelenid spiders of Turkiye and the Caucasus
are provided. Five species are described as new to science: Maimuna antalyensis sp. nov.
(39; Turkiye: Antalya), Tegenaria ballarini sp. nov. (49; Turkiye: Antalya), T. beyazcika
sp. nov. (3; Turkiye: Antalya), T. egrisiana sp. nov. (49; Georgia: Imereti), and T. hoeferi
sp. nov. (<4 9; Armenia: Kotayk). Tegenaria lazarovi Dimitrov, 2020, syn. nov. is proposed
as a new junior synonym of T. averni Brignoli, 1978. Persiscape caucasica (Gusein-
ov, Marusik & Koponen, 2005) is newly reported from Armenia, and T. chumachenkoi
Kovblyuk & Ponomarev, 2008 is reported for the first time from Turkiye. New distribution
records for T. dalmatica Kulczynski, 1906, T. hamid Brignoli, 1978, T. longimana Simon,
1898 and T. percuriosa Brignoli, 1972, and topotype material for T. tekke Brignoli, 1978
are reported. The record of Eratigena fuesslini (Pavesi, 1873) from Turkiye is found to
be based on a misidentification, and is herein attributed to T. hamid. The presence of an
embolic spine, unknown in any other species of Tegenaria, is documented in T. anhela
Brignoli, 1972 for the first time. Photographs are provided for all treated species.

Key words: Anatolia, Armenia, Georgia, Maimuna, new record, new synonymy, Persiscape,
Tegenaria

Introduction

Agelenidae C.L. Koch, 1837 is a large family of spiders, encompassing 1,405
extant species across 96 genera worldwide (WSC 2024). Commonly known as
“funnel-weavers, the family has been relatively well-studied in the Palaearc-
tic (e.g., de Blauwe 1980; Levy 1996). In the Western Palaearctic, Turkiye has
the highest recorded diversity of Agelenidae, with 74 species documented
(Danigman et al. 2024). Other areas within the Western Palaearctic remain
largely under-studied. For example, in the Caucasus, Otto (2022) lists 36 spe-
cies in ten genera of Agelenidae, yet only three species have been reported
from Armenia and 18 from Georgia to date. This highlights the limited under-
standing of agelenid diversity in this region (Zarikian et al. 2022).

291

Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

While examining spiders from Turkiye, Georgia, and Armenia, we had the
opportunity to study several agelenid specimens from these countries. In this
paper, we present the following findings: the descriptions of four new species
of Tegenaria Latreille, 1804 and of one new species of Maimuna Lehtinen, 1967;
the synonymization of T. /azarovi Dimitrov, 2020; the presence of an embolic
spine in T. anhela Brignoli, 1972; and several new faunistic data for agelenids
in Turkiye and Armenia.

Materials and methods

Photographs of specimens and their copulatory organs were obtained using
an Olympus Camedia E-520 camera attached to an Olympus SZX16 stereo-
microscope at the Zoological Museum of the University of Turku, Finland.
Digital images of different focal planes were stacked with Helicon Focus™
8.1.1. Illustrations of vulvae were made after digesting tissues off in a 10%
KOH aqueous solution. Body measurements exclude the chelicerae and spin-
nerets. Leg segments were measured on the dorsal side. Measurements of
legs are listed as: total (femur, patella, tibia, metatarsus, tarsus). All measure-
ments are given in millimeters.

Abbreviations used in the text and figures

Eyes: ALE—anterior lateral eye, AME—anterior median eye, PLE—posterior later-
al eye, PME—posterior median eye.

Leg segments: Fe—femur, Pa—patella, Ta—tarsus, Ti-— tibia.

Male palp: Eba—embolus base, Em—embolus, Es—embolic spine, Ce—claw-
like projections of the conductor, Cn—conductor, Cp—basal process of the
cymbium, Ma—median apophysis, Mp—median process, Rd-—retrodorsal tibial
apophysis, Rl—retrolateral tibial apophysis, Rv—retroventral tibial apophysis,
Ts—tooth of the retrolateral tibial apophysis, Ve—retroventral crest.

Epigyne: Cd—copulatory duct, Co—copulatory opening, Fo—fovea, Mr—mem-
branous part of the receptacle, Rs—sclerotized part of the receptacle, SIl—lon-
gitudinal scuta.

Depositories: MHNG—Muséum drhistoire naturelle, Genéve, Switzerland (L.
Monod); ZMUT—Zoological Museum of the University of Turku, Finland (V.
Vahtera); ZMMU—Zoological Museum of the Moscow State University, Russia
(K.G. Mikhailov); ZMUU—Zoological Museum of the Bursa Uludag University,
Turkiye (R.S. Kaya).

Taxonomy

Family Agelenidae C.L. Koch, 1837
Subfamily Ageleninae C.L. Koch, 1837

Tribe Agelenini C.L. Koch, 1837

Comment. For the diagnosis and composition, see Lehtinen (1967), Bolzern et
al. (2013), and Zamani and Marusik (2020).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Persiscape caucasica (Guseinov, Marusik & Koponen, 2005)
Figs 1C, 2D, E

Agelescape caucasica Guseinov et al., 2005: 157, figs 9-12, 69-71, 105 (9).
Persiscape caucasica: Zamani and Marusik 2020: 376, fig. 12M-O (9).

Material. ARMENIA: Kotayk Prov.: * 1 2 (ZMUT), env. of Geghadir, 40°09'N,
44°38'E, 15.05.2021 (Y.M. Marusik).

Comment. The male of this species is currently unknown.

Distribution. Previously known from Greece, Turkiye, Georgia, and Azerbaijan
(WSC 2024). A new record for Armenia.

Tribe Textricini Lehtinen, 1967
Comment. For the diagnosis and composition, see Kaya et al. (2023).

Maimuna antalyensis sp. nov.
https://Zzoobank.org/3A871504-2F88-4DF2-9053-90DFF4238122
Figs 1A, B, 2A-C, 3A-D, 4A-D

Type material. Holotype « 4 (ZMUT), TuRKIYE: Antalya Prov.: Alanya, env. Kestel,
Dim Valley, 36°32'34.5"N, 32°06'17.5"E, 110 m, pine and oak forest, 2.01.2013
(Y.M. Marusik). Paratypes: * 5 2 (ZMUT), same data as for the holotype;
*1 2 (ZMUT), Asmaca, 36°36'32.3"N, 32°03'12.4"E, 686 m, pine and oak forest,
3.01.2013 (Y.M. Marusik); «2 9 (ZMUT), Elikesik rd., 36°33'55.6"N, 31°55'30.3'E,
24 m, maquis on S exposed slope, 8.01.2013 (Y.M. Marusik); * 2 2 (ZMUT),
slopes of Alanya Castle, Damlatas side, 36°32'11.6"N, 31°59'30.3"E, 50 m, pine
forest, under stones and in litter, 7.01.2013 (Y.M. Marusik).

Comparative material. Maimuna vestita (C.L. Koch, 1841): TURKIYE: Bursa
Prov.: * 1 9 (ZMUU), Bursa Uludag University campus area, 30.11.1999 (R.S.
Kaya); * 2 9 (ZMUU), same, 16.05.2000 (R.S. Kaya); * 1 9 (ZMUU), same,
3.03.2003 (R.S. Kaya); * 1 2 (ZMUU), same, 10.09.2005 (R.S. Kaya); 1419
(ZMUU), same, 20.04.2012 (R.S. Kaya); * 1 3 1 2 (ZMUU), same, 21.05.2012
(R.S. Kaya); * 5 9 (ZMUU), same, 4.05.2023 (R.S. Kaya); * 2 3 3 2 (ZMUU),
same, 4.01.2024 (R.S. Kaya); * 1 4 2 9 (ZMUU), same, 11.07.2024 (R.S. Kaya);
- 5 2 (ZMUU), Lake Uluabat, Halilbey Island, 8.04.2001 (R.S. Kaya);* 143 9
(ZMUU), Lake Uluabat, Terzioglu Island, 14.10.2004 (R.S. Kaya); * 5 2 (ZMUU),
same, 25.04.2005 (R.S. Kaya); * 1 3 2 9 (ZMUU), Lake Uluabat, Manastir Island,
28.09.2005 (R.S. Kaya); * 1 4 (ZMUU), same, 29.09.2005 (R.S. Kaya);* 2419
(ZMUU), Lake Uluabat, Halilbey Island, 15.12.2005 (R.S. Kaya); * 1 9 (ZMUU),
Karacabey, Bogaz, 4.05.2005 (R.S. Kaya); » 3 2 (ZMUU), same, 8.08.2007
(R.S. Kaya); * 2 2 (ZMUU), same, 5.06.2018 (R.S. Kaya); * 2 2 (ZMUU), same,
11.06.2021 (R.S. Kaya); + 1 ¢ 1 2 (ZMUU), Kaplikaya, 29.03.2007 (R.S. Kaya);
- 2 3 2 9 (ZMUU), same, 10.12.2008 (R.S. Kaya); * 1 2 (ZMUU), Goriikle Vill.,
20.04.2012 (R.S. Kaya); * 1 2 (ZMUU), Niliifer Metro station, 26.05.2012 (R.S.
Kaya); * 1 4 (ZMUU), Orhangazi Dist., 4.05.2014 (R.S. Kaya); Canakkale Prov.:
* 1 2 (ZMUU), Gokceada Island, Lake salt area, 4.05.2004 (R.S. Kaya).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

imm

rer
—

We ‘

Figure 1. Habitus of Maimuna antalyensis sp. nov. (A, B), Persiscape caucasica (C), and Tegenaria hamid (D), dorsal view.
A male B-D females.

Diagnosis. The new species is similar to M. cariae Brignoli, 1978 in the over-
all shape of its copulatory organs. The male differs by having a shorter tip of
the cymbium (as long as the palpal tibia, vs longer), and by a different shape of
the conductor and the median process (cf. Fig. 3B-D and Dimitrov 2022: fig.
26). The female of the new species has a hexagonal epigynal fovea, in contrast
to the subtriangular fovea of M. cariae (cf. Fig. 2A, B and Dimitrov 2022: fig. 17).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

bv Se :
Figure 2. Epigyne of Maimuna antalyensis sp. nov. (A-C) and Persiscape caucasica (D, E). A macerated, ventral view
B, D vulva, dorsal view C, E intact, ventral view. Abbreviations: Co — copulatory opening, Fo — fovea. Scale bars: 0.2 mm.

Pe.

Description. Male. Habitus as in Fig. 1A. Total length 5.40. Carapace 2.60
long, 1.90 wide. Eye sizes: AME: 0.10, ALE: 0.13, PME: 0.18, PLE: 0.13. Carapace,
sternum, labium, and maxillae pale brown; carapace with darker submedian
bands; ocular region black. Legs yellowish brown, with annulations. Abdomen
dorsally dark greyish with paler chevrons, pale greyish ventrally. Spinnerets pale
greyish, darker basally. Measurements of legs: |: 5.81+missing Ta (1.74, 0.82,
1.43, 1.82, missing), Il: 6.73 (1.77, 0.82, 1.37, 1.73, 1.04), Ill: 6.64 (1.74, 0.76,
1.34, 1.74, 1.06), IV: 8.53 (2.19, 0.85, 1.80, 2.55, 1.14).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

Figure 3. Male palp of Maimuna antalyensis sp. nov. A ventral view B retroventral view C femur, ventral view D retrolateral
view. Abbreviations: Eb — base of the embolus, Em - embolus, Cn — conductor, Mp - median process, Vc — retroventral
crest of the tibia. Scale bars: 0.2 mm.

Palp as in Figs 3A-D, 4A-D; femur modified — with retroventral bump in mid
part, ~ 3x longer than wide, shorter than cymbium; dorsal length of patella same
as in tibia; tibia ~ 1.4x wider than long (Fig. 3C, D), lacking prominent apophysis
but with retroventral crest (Vc) (Fig. 3A, B); cymbium ~ 1.4x longer than wide, with
tip ~ 1/3 of cymbial length; bulb as long as wide; conductor (Cn) massive, strong-
ly sclerotized prolateral arm lacking, terminal part with two claw-like projections
(Cc; Fig. 4D); median process (Mp) massive, roundly bent in lateral view (Fig. 4B,
C); embolus (Em) filiform, weakly sclerotized, with small base (Eb).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

Figure 4. Dissected bulb of Maimuna antalyensis sp. nov. A proximal view B ventral view C proventral view D retroventral
view. Abbreviation: Cc — claw-like projections of the conductor. Scale bars: 0.2 mm.

Female. Habitus as in Fig. 1B. Total length 7.50. Carapace 3.28 long, 2.07
wide. Eye sizes: AME: 0.12, ALE: 0.17, PME: 0.20, PLE: 0.15. Coloration as in
male. Measurements of legs: I: 7.25 (1.94, 1.02, 1.44, 1.76, 1.09), Il: 7.24 (1.85,
1.05, 1.47, 1.73, 1.14), Ill: 7.37 (2.00, 0.96, 1.42, 1.94, 1.05), IV: 9.58 (2.45, 1.11,
2.137266, 1:23).

Epigyne as in Fig. 2A—C; epigynal plate slightly wider than long; fovea (Fo)
hexagonal, approximately as long as wide, located anteriorly (Fig. 2C); copulatory
ducts gradually turning to receptacles and approximately as wide as receptacles;
copulatory ducts converging and contiguous, receptacles diverging (Fig. 2A, B).

Comment. In the examined comparative female specimens of M. vestita, we
observed noticeable variation in both body size and epigyne morphology. In
particular, in smaller individuals, the shape of the epigynal fovea can vary sig-
nificantly. A similar pattern is observed in M. antalyensis sp. nov., where smaller
females also exhibit variation in the shape of the epigynal fovea, ranging from
hexagonal to nearly circular in some specimens.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 5. Epigyne of Tegenaria hamid. A intact, ventral view B macerated, anteroventral view C vulva, dorsal view D vulva,
anterodorsal view. Scale bars: 0.2 mm.

Note. For comments on the homology of the structure referred to here as the
“median process,” see Kaya et al. (2023).

Distribution. Known from the listed localities in Antalya Province, southwest-
ern Turkiye.

Etymology. The specific epithet refers to the type locality of the species in
Antalya, Turkiye.

Tribe Tegenariini Lehtinen, 1967
Comment. For the diagnosis and composition, see Bolzern et al. (2010).

Tegenaria anhela Brignoli, 1972
Figs 6A, B, 7A-D, 8A-C, 20B

Tegenaria anhela Brignoli, 1972: 173, figs 24-27 (¢9).
Malthonica anchela: Guseinov et al. 2005: 164 (lapsus).
Tegenaria anhela: Bolzern et al. 2013: 846.

Material. TURKIYE: Antalya Prov.: * 1 4 1 2 (ZMUT), Désemealti, 37°01'N,
30°36'E, 4.08.2009 (R.S. Kaya, C. Kaya); + 1 ¢ 1 2 (ZMUU). same data; * 2 9
(ZMUU), Désemealti, Karain Cave (R.S. Kaya, C. Kaya).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

1mm

Figure 6. Habitus of Tegenaria anhela (A, B) and Tegenaria ballarini sp. nov. (C, D). A, C females B, D males.

Comments. Tegenaria anhela was described based on material collected
from Karain Cave in Antalya (Brignoli 1972). Later, it was recorded from Mustan
Ini Cave, which is located near the type locality (Brignoli 1978a). Although the
original description includes informative illustrations, it lacks detailed expla-
nations of the palpal structures. During the examination of our material, we

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

Figure 7. Male palp of Tegenaria anhela. A ventral view B retrolateral view C prolateral view D dorsal view. Abbreviation:
Es — embolic spine. Scale bars: 0.2 mm.

observed a unique structure, which we term the “embolic spine” (Es; Fig. 7A).
This structure, the homology of which remains unclear, is situated between the
base of the embolus and the conductor, and it has not been reported in any
other Tegenaria species to date.

Distribution. Known only from Antalya Province, southwestern Turkiye.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Tegenaria averni Brignoli, 1978
Figs 8D, E, 13A-D

Tegenaria averni Brignoli, 1978a: 50, fig. 10 (@).
Tegenaria lazarovi Dimitrov, 2020: 48, figs 1-12 (42). Syn. nov.

Material. TuRKIYE: Mersin Prov.: * 2 3 2 2 (ZMUT), Silifke, Cennet Cave,
36°26'12'N, 34°06'22"E, 20.09.2010 (Y.M. Marusik).

Comments. Tegenaria averni was described based on a single female from
Cennet Cave in Mersin (Brignoli 1978a). In his review of the ariadnae spe-
cies-group, Dimitrov (2020) described T. lazarovi based on material of both
sexes collected in a cave in Mersin. Although the type locality of T. lazarovi
is only 65 km away from that of T. averni, and both are in the same provincial
district, T. averni is not mentioned in Dimitrov (2020) at all, likely due to a lack
of males. Collection of material of both sexes at the type locality of T. averni
revealed that these two populations are conspecific. Therefore, T. lazarovi syn.
nov. is proposed as a junior synonym of T. averni.

Distribution. Known only from two caves in Mersin Province, southern Turkiye.

Tegenaria ballarini sp. nov.
https://Zzoobank.org/BB2AE92D-4A0E-4BFD-9FC5-8F 2434070636
Figs 6C, D, 9A-D, 10A, B, 12A-C

Type material. Holotype « ¢ (ZMUU), TuRKIYE: Antalya Prov.: Bozyaka, K6prilii
Canyon National Park, 37°11'51"N, 31°11'03"E, 243 m, 15.05.2008 (R.S. Kaya).
Paratypes: +1 3 2 2 (ZMUT), same data as for the holotype; * 7 2 (ZMUU), same
data as for the holotype.

Comparative material. Tegenaria vankeerorum Bolzern, Burckhardt & Hang-
gi, 2013 (Figs 10C, D, 11A—D): TURKIYE: Mugla Prov.: * 1 41 9 (ZMUT), Yatagan,
Orman Isletme, 37°20'N, 28°08'E, 18.05.2011 (R.S. Kaya).

Diagnosis. The new species is closely related to T. vankeerorum and has very
similar copulatory organs, especially the male palp. The male of T. ballarini sp.
nov. differs from the similar species by having relatively longer palpal tibia and a
retrolateral apophysis (R/) located in the distal half of the tibia, rather than at the
midpoint (cf. Figs 9C, 10B, 11B). The female of the new species differs from all
other species of Tegenaria by having a pair of longitudinal scuta (S/) anterior to
the epigynal plate and a straight posterior margin of the epigyne (Fig. 12C). Addi-
tionally, the vulva of the new species differs from that of T. vankeerorum by having
relatively longer copulatory ducts that almost reach the anterior margin of the
receptacle (vs reaching only the mid part of the receptacle; cf. Figs 12A, B, 10D, E).

Description. Male. Habitus as in Fig. 6D. Total length 10.75. Carapace 6.75
long, 4.65 wide. Eye sizes: AME: 0.25, ALE: 0.25, PME: 0.22, PLE: 0.25. Pars ce-
phalica, sternum, labium, maxillae, and Fe | brown; pars thoracica and remain-
ing leg segments yellowish brown; chelicerae dark reddish brown. Legs with-
out annulations. Abdomen dark grey, without patterns. Spinnerets uniformly
greyish brown. Measurements of legs: |: 48.87+missing Ta (14.28, 2.82, 14.60,
17.17, missing), Il: 44.00 (12.00, 2.72, 11.50, 14.28, 3.50), Ill: 36.83 (9.98, 2.45,
8.65, 12.65, 3.10), IV: 44.36 (11.56, 2.50, 10.88, 15.82, 3.60).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 8. Epigyne of Tegenaria anhela (A-C) and T. averni (D, E). A, D macerated, ventral view B, E vulva, dorsal view
C intact, ventral view. Scale bars: 0.2 mm.

Palp as in Figs 9A-D, 10A, B; femur 5x longer than wide and ~ 1.8x longer
than tibia; patella ~ 1.9x longer than wide; tibia long, ~ 3.5x longer than wide,
with three apophyses: retrolateral (R/), retroventral (Rv) and retrodorsal (Rd)
(Fig. 9A-C); retrolateral apophysis located in distal 1/3 of tibia, spine-like, di-
rected antero-retrolaterally, with small tooth (Ts) (Fig. 9B); cymbium long, 2.6x
longer than wide, tip as long as bulb (Fig. 10B); bulb as long as wide (account-
ing conductor), tegulum oval, bent prolaterally, and distal part extending em-
bolus; conductor (Cn) with rounded distal arm and gradually tapering proximal
arm with claw-like tip, terminated at ~ 3 o'clock position; embolus originates at
9 o'clock position, thick basally, roundly bent (Fig. 10A).

Female. Habitus as in Fig. 6C. Total length 14.15. Carapace 8.20 long, 6.15
wide. Eye sizes: AME: 0.25, ALE: 0.27, PME: 0.28, PLE: 0.31. Coloration as in
male. Measurements of legs: |: 42.17 (11.61, 3.40, 11.17, 12.34, 3.65), Il: 37.29
(10.48, 2.98, 9.36, 11.12, 3.35), Ill: 33.05 (9.40, 2.75, 7.70, 10.20, 3.00), IV: Fe:
11.03, Pa: 2.98, Ti: 10.03, remaining segments missing.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Tu SO

Figure 9. Male palp of Tegenaria ballarini sp. nov. A ventral view B dorsal view C retrolateral view D prolateral view. Abbre-
viations: Cn — conductor, Rd — retrodorsal apophysis, R/ — retrolateral apophysis, Rv — retroventral apophysis, Ts — small
tooth of the retrolateral apophysis.

Epigyne as in Fig. 12A—C; epigynal plate > 2x wider than long, with straight
and heavily sclerotized posterior margin; area anteriorly from plate with pair of
elongate scuta (S/) (Fig. 12C); receptacles subdivided into two parts, anterior
pear-shaped membranous part (Mr) and heavily sclerotized posterior parts (Rs)
(Fig. 12B); copulatory ducts well sclerotized, thin, contiguous, terminating near
anterior edges of receptacles.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 10. Copulatory organs of Tegenaria ballarini sp. nov. (A, B) and T. vankeerorum (C-E). A, C palp, ventral view B full
palp, retrolateral view D vulva, dorsal view E macerated epigyne, ventral view. Abbreviation: Cn — conductor. Scale bars:
0.2 mm, unless otherwise indicated.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

0.5mm

WwW G0

0.5mm

WW G0

Figure 11. Male palp of Tegenaria vankeerorum. A ventral view B retrolateral view C dorsal view D prolateral view.

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Figure 12. Epigyne of Tegenaria ballarini sp. nov. A macerated, ventral view B vulva, dorsal view C intact, ventral view.
Abbreviations: Mr — membranous part of the receptacle, Rs — sclerotized part of the receptacle, S/ — longitudinal scutum
of the epigynal plate. Scale bars: 0.2 mm, unless otherwise indicated.

Distribution. Known only from the type locality in Antalya Province, south-
western Turkiye.

Etymology. The new species is named in honor of our colleague Francesco
Ballarin (Tokyo, Japan), in recognition of his assistance to the second author
during her visit to the Brignoli collection in Verona, Italy.

Tegenaria beyazcika sp. nov.
https://zoobank.org/2EEF98AE-7348-4DEA-AE26-AEO50EFBA76A
Figs 14A-D, 18D

Type material. Holotype « 4 (ZMUT), TuRKIYE: Antalya Prov.: Alanya, env. Kestel,
Dim Valley, 36°32'34.5"N, 32°06'17.5"E, 110 m, pine and oak forest, 2.01.2013
(Y.M. Marusik). Paratypes: * 4 ¢ (ZMUT), same data as for the holotype.

Diagnosis. The new species belongs to the ariadnae species-group and is most
similar to T. averni. The male of the new species differs from that of T. averni by
having thickened male palpal femur with four strong dorsal spines (Fig. 14B), an
almost straight embolus on the prolateral half (vs roundly bent), a relatively shorter
tibia with a length/width ratio of 2.5 (vs 2.9), and a conductor with subequal arms
(vs a distal arm that is longer than the proximal arm; cf. Figs 14A, 13A).

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Figure 13. Male palp of Tegenaria averni. A ventral view B dorsal view C prolateral view D retrolateral view. Scale bars: 0.2 mm.

Description. Male. Habitus as in Fig. 18D. Total length 4.00. Carapace 1.95
long, 1.55 wide. Eye sizes: AME: 0.04, ALE: 0.07, PME: 0.06, PLE: 0.07. Pars ce-
phalica, chelicerae, labium, maxillae, and Fe | and II pale brown, Fe II paler than |;
pars thoracica, sternum, and remaining leg segments pale brown. Legs without
annulations. Fe |, and to lesser degree Fe II, with ventral coating of long setae.
Abdomen pale beige, without patterns. Spinnerets uniformly pale beige. Measure-
ments of legs: I: 9.00 (2.36, 0.78, 2.24, 2.24, 1.38), Il: 7.95 (2.16, 0.74, 1.85, 1.93,
1.27), Ill: 7.39 (1.93, 0.67, 1.65, 2.00, 1.14), IV: 9.78 (2.55, 0.75, 2.36, 2.74, 1.38).

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Figure 14. Male palp of Tegenaria beyazcika sp. nov. A ventral view B full palp, retrolateral view C prolateral view D retro-
lateral view. Abbreviations: Ma - median apophysis, Rd — retrodorsal apophysis. Scale bars: 0.2 mm.

Palp as in Fig. 14A—D; femur 4x longer than wide, longer than cymbium, 1.5x
wider than tibia, with 4 strong spines in distal 1/2 (Fig. 14B); patella 2x longer
than wide; tibia 2.25x longer than wide, with retroventral (Rv) and retrodorsal
(Rd) apophyses (Fig. 14B); cymbium > 2x longer than wide, tip approximately
as long as cymbium wide, with two strong macrosetae (= spines) on retrolateral

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1/2; bulb as long as wide; median apophysis (Ma) long, approximately as long
as width of tibia, originating at ~ 5 o’clock position (Fig. 14A); conductor fun-
giform, with both arms of equal length and width; embolus originating at ~
8 o'clock position, straight in prolateral 1/2 of bulb and strongly roundly bent
proximally at retrolateral side.

Female. Unknown.

Comments. Although the specimens of both T. hamid and T. beyazcika
sp. nov. (known only from females and males, respectively) were collected
from the same locality, we consider them to belong to different species due
to noticeable differences in size and coloration. Additionally, T. hamid has
a different conformation of the copulatory organs compared to those of the
species in the ariadnae group, thus belonging to a different species-group
than T. beyazcika sp. nov. Given the pale coloration of this species, the rela-
tively elongated legs, and the dense ventral coating of long setae on femora
| and Il, it seems that the collection locality mentioned on the label is slightly
off. It is more likely that the species was collected from a cave, such as the
nearby Dim Cave.

Distribution. Known only from the type locality in Antalya Province, south-
western Turkiye.

Etymology. The specific epithet is derived from the Turkish word "beyaz’",
meaning pale, combined with the suffix -cik, meaning little. This refers to the
relatively small size and pale coloration of this species.

Tegenaria chumachenkoi Kovblyuk & Ponomarev, 2008
Fig. 15A-D

Tegenaria chumachenkoi Kovblyuk & Ponomarev, 2008: 147, figs 18-21 (9).
Tegenaria chumachenkoi: Ponomarev and Shmatko 2022: 212, figs 5-10 (32).
— Seropian et al. 2023: 147, suppl.: 5 fig. (3).

Material. TURKIYE: Artvin Prov.: * 1 3 (ZMUT), Savsat Dist., env. of Meydancik
Town, Erikli Vill.,41°27'13.1"N, 42°13'23.8"E, 1141 m, 12.06.2009 (Y.M. Marusik).

Distribution. Previously known from Azerbaijan, Georgia, and northern Cau-
casus (Ponomarev and Shmatko 2022). A new record for Turkiye.

Tegenaria dalmatica Kulczynski, 1906
Figs 16A-C, 18C

Malthonica dalmatica: Kovblyuk and Nadolny 2007: 19, figs 1-10 (¢).
Tegenaria dalmatica: Bolzern et al. 2013: 793, figs 1G, H, 2A, B, D, 15K, L, O, Q (39).

Note. For a full list of 14 taxonomic entries, see WSC (2024).

Material. TURKIYE: Izmir Prov.: * 6 2 (ZMUT), Kemalpasa, Visneli Vill., Fetrek-2
Cave, 38°20'N, 27°25'E, 311 m, 5.06.2009 (Y.M. Marusik); Bursa Prov.: * 1 3
(ZMUU), Lake Uluabat, Terzioglu Island, 21.11.2003 (R.S. Kaya); * 1 9 (ZMUU),
Lake iznik, Gélliice Vill., 14.10.2016, (R.S. Kaya).

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Figure 15. Male palp of Tegenaria chumachenkoi. A ventral view B retrolateral view C dorsal view D prolateral view.
Abbreviation: Ma - median apophysis. Scale bars: 0.2 mm.

Comment. The only previous record of this species from Turkiye was by
Bolzern et al. (2013), although lacking further locality data.

Distribution. From Iberian Peninsula to Turkiye, south to northern Africa
(Nentwig et al. 2024; WSC 2024).

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Figure 16. Epigyne of Tegenaria dalmatica (A-C) and T. egrisiana sp. nov. (D-F). A, E macerated, ventral view B, F vulva,
dorsal view C, D intact, ventral view. Abbreviations: Cd — copulatory duct, Co — copulatory opening. Scale bars: 0.2 mm.

Tegenaria egrisiana sp. nov.
https://zoobank.org/45CD9A9F-8B0B-47DD-B0A7-145458C6B6BE
Figs 16D-F, 17A-D, 18A, B, 20C

Type material. Holotype + ¢ (ZMMU), GEorGiA: Imereti Prov.: cave between Gum-
brini and Khamali, 42°18'56.4"N, 42°38'09.4"E, 161 m, 19.07.2012 (Y.M. Marusik).
Paratypes: +1 31 9 (ZMUT), 2 2 (ZMMU), same data as for the holotype.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 17. Male palp of Tegenaria egrisiana sp. nov. A ventral view B prolateral view C dorsal view D retrolateral view.
Abbreviations: Ma - median apophysis, RI — retrolateral apophysis, Rv — retroventral apophysis. Scale bars: 0.2 mm.

Diagnosis. Tegenaria egrisiana sp. nov. is very similar to T. pallens Zamani &
Marusik, 2023 from Iran in the overall shape of the copulatory organs. Howev-
er, the male differs from T. pallens in the shorter tip of the cymbium, ~ 0.7 the
length of the palpal tibia (Fig. 17A—D, 20C; vs as long as the palpal tibia), the

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 18. Habitus of Tegenaria egrisiana sp. nov. (A, B), T. dalmatica (C), T. beyazcika sp. nov. (D), and T. tekke (E), dorsal
view. A, D males B, C, E females.

blunt tip of the conductor (vs pointed and curved; Zamani et al. 2023: fig. 2A),
the embolus base positioned at the 9:00 o'clock position (vs 8:30 o'clock), the
tip of the embolus terminating at ~ 2:00 o'clock position (Fig. 17A; vs 1:00
o'clock), and the median apophysis (Ma) with a different shape. The female of

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 19. Male palp of Tegenaria longimana. A ventral view B prolateral view C dorsal view D retrolateral view. Scale
bars: 0.2 mm.

the new species differs from that of T. pallens in the epigynal plate nearly twice
as wide as it is long (vs > 3x wider than long; cf. Fig. 16D and Zamani et al.
2023: fig. 3C), in having a distinct median plate (vs absent), and a small rectan-
gular fovea (vs oval; cf. Fig. 16D, E and Zamani et al. 2023: fig. 3A, B).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

Figure 20. Full male palp of Tegenaria longimana (A), T. anhela (B), and T. egrisiana sp. nov. (C). A prolateral view

B, C retrolateral view.

Description. Male. Habitus as in Fig. 18A. Total length 7.10. Carapace 3.23
long, 2.37 wide. Eye sizes: AME: 0.12, ALE: 0.17, PME: 0.14, PLE: 0.18. Cara-
pace, labium, and maxillae pale brown; carapace with darker submedian bands;
chelicerae reddish brown; sternum greyish brown, with yellow median band and
six spots. Legs pale brown, with very faint annulations; Fe with long ventral se-
tae at basal 1/2. Abdomen pale beige, with greyish dots, patches, and stripes.
Spinnerets uniformly pale beige. Measurements of legs: I: 22.92 (6.12, 1.44,
6.26, 6.50, 2.60), Il: 19.02 (5.05, 1.28, 4.96, 5.50, 2.23), Ill: 16.90 (4.63, 1.15,
4.15, 5.15, 1.82), IV: 20.79 (5.50, 1.28, 5.24, 6.65, 2.12).

Palp as in Fig. 17A—D; femur longer than patella+tibia; femur ~ 2.2x longer than
tibia (Fig. 20C); patella 2x longer than wide; cymbium ~ 1.8x longer than tibia; tib-
ia ~ 2x longer than wide, with two apophyses: large and membranous retroven-
tral apophysis (Rv) and conical retrolateral apophysis (R/) with a notched blunt tip
(Figs 17D, 20C); retrolateral apophysis shorter than ventrolateral one; cymbium 2x
longer than wide; bulb longer than wide; median apophysis (Ma) large and wide,
originating at ~ 4 oclock position; conductor as long as wide, with a spatula-like
tip; embolus filiform, roundly bent, originating at ~ 9:00 o'clock position (Fig. 17A).

Female. Habitus as in Fig. 18B. Total length 8.68. Carapace 4.25 long, 2.95
wide. Eye sizes: AME: 0.12, ALE: 0.20, PME: 0.18, PLE: 0.20. Coloration as in
male. Measurements of legs: |: 22.80 (6.13, 1.65, 6.00, 6.30, 2.72), Il: 19.81
(5:53, 1.57, 4.96, 5:47; 2.28), Ile 18:03 (5:07, 444,4°32,-5.22,-1.98), Vi 22.57
(6.18, 1.47, 5.63, 7.00, 2.29).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

A

Figure 21. Male palp of Tegenaria percuriosa. A ventral view B prolateral view C dorsal view D retrolateral view. Scale
bars: 0.2 mm.

Epigyne as in Fig. 16D-F; epigynal plate ~ 2x wider than long with two scle-
rotized and barely visible teeth; fovea small and almost rectangular; copulatory
openings (Co) located on anterior edges of holes (Fig. 16D, E); copulatory ducts
(Cd) with a membranous anterior part and a widened slightly sclerotized poste-
rior part; receptacles tubular and twisted along their axis (Fig. 16F).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 22. Epigyne of Tegenaria percuriosa (A, B) and T. longimana (C, D). A, C macerated, ventral view B, D vulva, dorsal

view. Scale bars: 0.2 mm.

Distribution. Known only from the type locality in Imereti Province, cen-
tral-western Georgia.

Etymology. The specific epithet refers to the historical Georgian polity of
Egrisi, which was centered in present-day western Georgia.

Tegenaria hamid Brignoli, 1978
Figs 1D, 5A-D

Tegenaria hamid Brignoli, 1978b: 515, fig. 96 (2).
Eratigena fuesslini: Topgcu and Demircan 2018: 20, fig. 1 (9).

Type material. Holotype * 2 (MHNG), TuRKIvE: Isparta Prov.: Egridir, 18.04.1973
(P. Brignoli). [examined]

Other material. TURKIYE: Antalya Prov.: * 2 9 (ZMUT), Alanya, env. Kestel,
Dim Valley, 36°32'34.5"N, 32°06'17.5"E, pine and oak forest, 2—9.01.2013 (Y.M.
Marusik); * 1 9 (ZMUT), Asmaca, 36°36'32.3"N, 32°03'12.4"E, 686 m, pine and
oak forest, 3.01.2013 (Y.M. Marusik).

Comment. This species was previously known only from its original de-
scription. The single figure of the vulva provided by Brignoli (1978b) is rather

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

schematic and not depicted from an exact dorsal view. Therefore, we present
additional figures from various angles based on newly collected material from
Antalya. Additionally, upon checking the record of Eratigena fuesslini (Pavesi,
1873) from Turkiye by Topcu and Demircan (2018), it became evident that the
figure presented corresponds to the anteroventral view of the macerated epi-
gyne of T. hamid, rather than E. fuesslini. Consequently, the record of E. fuesslini
is hereby removed from the list of Turkish spiders.

Distribution. Known only from Isparta and Antalya provinces, southwestern
Turkiye.

Tegenaria hoeferi sp. nov.
https://zoobank.org/3606E8DA-51F 1-4AB9-81 F1-6354AAA1456C
Figs 23A-D, 24A-C, 25A-C

Type material. Holotype > ¢ (ZMUT), ARMENIA: Kotayk Prov.: env. Aghveran,
40°29'54"N, 44°35'24"E, 7-8.05.2021 (Y.M. Marusik). Paratypes: > 1 3 3 9
(ZMUT, ZMMU), same data as for the holotype.

Diagnosis. The new species belongs to the abchasica species-group and is
most similar to T. chumachenkoi. The male of the new species differs from that
of T. chumachenkoi by the shape of the median apophysis, bulging proximally
and widely pointed retrolaterally (vs straight proximally and sharply pointed ret-
rolaterally in T. chumachenkoi; cf. Figs 23A, 15A). The female of the new spe-
cies differs from that of T. chumachenkoi by having an oval median plate that is
~ 2x as wide as it is long (vs the median plate is not oval and is approximately
as long as it is wide; cf. Fig. 25A and Ponomarev and Shmatko 2022: fig. 10).

Description. Male. Habitus as in Fig. 24B. Total length 8.40. Carapace 4.10
long, 3.20 wide. Eye sizes: AME: 0.20, ALE: 0.21, PME: 0.17, PLE: 0.19. Cara-
pace, chelicerae, labium, and maxillae pale brown; carapace with black subme-
dian and marginal bands; sternum greyish brown, with yellow median lobulated
band and six spots. Legs pale brown, with distinct annulations. Abdomen dark
greyish, with numerous beige dots, patches, and stripes. Anterior spinnerets
greyish basally and pale beige apically, posterior ones uniformly pale beige.
Measurements of legs: |: 17.48 (4.56, 1.62, 4.06, 4.91, 2.33), Il: 17.14 (4.40,
1.63, 4.07, 4.71, 2.33), Ill: 15.86 (4.29, 1.47, 3.57, 4.48, 2.05), IV: 19.60 (5.16,
1.52, 4.58, 6.08, 2.26).

Palp as in Figs 23A-D, 24C; femur roundly bent, ~ 4x longer than wide; pa-
tella swollen, approximately as wide as long with long dorsal seta almost as
long as tibia; tibia ~ 1/2 as long as femur (not counting apophyses) (Fig. 24C);
retrodorsal apophysis (Rd) approximately as long as tibia wide distally, with
strong spine directed proximoventrally (Fig. 23B); retroventral apophysis (Rv)
small (Fig. 23C, D); cymbium 1.8x longer than wide, tip ~ 1/3 of cymbial length,
lacking distinct spine, with basal process (Cp); bulb as long as wide; median
apophysis (Ma) large (Fig. 23A), bent prolaterally; conductor with long and thin
distal arm not reaching prolateral 1/2 of cymbium; embolus with large base,
free part originating at 9 o'clock position, thin, roundly bent.

Female. Habitus as in Fig. 24A. Total length 10.28. Carapace 4.00 long, 3.00
wide. Eye sizes: AME: 0.18, ALE: 0.20, PME: 0.16, PLE: 0.18. Coloration as in
male. Measurements of legs: |: 16.95 (4.33, 1.69, 4.18, 4.55, 2.20), Il: 15.92

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Figure 23. Male palp of Tegenaria hoeferi sp. nov. A ventral view B retrolateral view C dorsal view D prolateral view.
Abbreviations: Cp — basal process of the cymbium, Rd - retrodorsal apophysis, R/ — retrolateral apophysis, Rv — retro-
ventral apophysis. Scale bars: 0.2 mm.

(4.20, 1.63, 3.65, 4.15, 2.29), Ill: 14.43 (3.94, 1.43, 3.23, 4.00, 1.83), IV: 17.89
(4.74, 1.58, 4.21, 5.29, 2.07).

Epigyne as in Fig. 25A-C; plate 2x wider than long, median plate oval, wider
than long (Fig. 25A, B); copulatory ducts and receptacles lacking distinct limits,
contiguous (Fig. 25C).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 24. Tegenaria hoeferi sp. nov. A, B habitus, dorsal view C full palp, retrolateral view. A female B, C male.

Distribution. Known only from the type locality in Kotayk Province, central
Armenia.

Etymology. This species is named after Hubert H6fer (Karlsruhe, Germany),
a German arachnologist. He is the Curator of Invertebrates and head of Biosci-
ences at the State Museum of Natural History Karlsruhe. He has made signifi-
cant contributions to the study of spiders in both South America and Germany,
leading numerous projects and helping to compile the largest dataset on distri-
butions of spiders in Germany.

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Tegenaria longimana Simon, 1898
Figs 19A-D, 20A, 22C, D

Tegenaria longimana: Ponomarev et al. 2024: 279, figs 29-33 (¢9).

Note. For a full list of seven taxonomic entries, see WSC (2024).

Material. GEORGIA: Imereti Prov.: > 1 4 4 9 (ZMUT), env. of Tskhaltubo,
Khomuli Vill., Tetra Cave, 42°19'49.3"N, 42°37'00.9"E, 454 m, 18.07.2012 (Y.M.
Marusik); * 1 2 (ZMUT), Bzvani Vill., deep cave, near the entrance, 42°03'01.3'N,
42°36'04.5"E, 402 m, 20.07.2012 (Y.M. Marusik).

Comment. This species was described from Batumi, the capital of the Geor-
gian republic of Adjara.

Distribution. Known from Turkiye, Georgia, and northern Caucasus (Otto
2022; WSC 2024).

Tegenaria percuriosa Brignoli, 1972
Figs 21A-—D, 22A, B

Tegenaria percuriosa: Dimitrov et al. 2022: 3, figs 1, 2, 7-13, 40, 41 (49).

Note. For a full list of five taxonomic entries, see WSC (2024).

Material. TURKIYE: Antalya Prov.: * 1 9 3j. (ZMUT), Alanya, env. Kestel,
Dim Cave, 36°32'22.1"N, 32°06'34.4"E, 225 m, 4.01.2013 (Y.M. Marusik);
Bursa Prov.: * 1 6. 1 9 (ZMUT), Uludag, G6yniikbelen rd., 39°59'N, 29°02'E,
14.05.2006 (R.S. Kaya); * 2 5 2 2 (ZMUT), InegGl, Great Oylat Cave, 39°56'N,
29°35'E, 519 m, 3.06.2009 (Y.M. Marusik); * 1 3 1 2 (ZMUU), Uludag Moun-
tain, Barakli Pond, 27.05.2003 (R.S. Kaya); * 1 4 (ZMUU), same, 8.05.2005
(R.S. Kaya); * 1 ¢ (ZMUU), same, 10.05.2010 (R.S. Kaya); * 1 4d (ZMUU), same,
1270 m, 16.05.2016 (R.S. Kaya); * 1 2 (ZMUU), Uludag Mountain, Soguk-
pinar Valley, 40°03'N, 29°09'E, 5.06.2003 (R.S. Kaya); * 1 2 (ZMUU), Uludag
Mountain, Alagam Forest, 30.08.2009 (R.S. Kaya); * 1 2 (ZMUU), Uludag
Mountain, Kocayayla Plateau, 25.09.2010 (R.S. Kaya); * 1 3 14 9 (ZMUU),
Uludag Mountain, National Park, 40°06'N, 29°05'E, 2.06.2010 (R.S. Kaya); * 1
& 2 9 (ZMUU), Kazanpinar Cave, 2.06.2009 (R.S. Kaya); * 2 4 10 2 (ZMUU),
same, 6.06.2009 (R.S. Kaya); * 2 ¢ 3 9 (ZMUU), Ayvaini Cave, 14.10.2012
(R.S. Kaya); * 1 2 (ZMUU), Oylat Cave, 15.10.2016 (R.S. Kaya); 2429
(ZMUU), Mustafakemalpasa Dist., Suuctu Waterfall, 24.06.2012 (R.S. Kaya);
Balikesir Prov.: * 1 2 (ZMUU), Alagam Mountain, 39°25'N, 38°35'E, 4.07.2012
(R.S. Kaya); Eskisehir Prov.: * 1 2 (ZMUU), Catacik Forest, 39°57'N, 31°08’E,
1.08.2012 (R.S. Kaya); Isparta Prov.: * 1 9 (ZMUU), Zindan Cave, 21.05.2007
(R.S. Kaya); Istanbul Prov.: > 1 2 (ZMUU), Aydos Forest, 40°56'N, 29°14'E,
874 m, 30.04.2016 (R.S. Kaya).

Comment. In our male specimens, the tip of the apical part of the median
apophysis is widened (Fig. 21A), which is different from that of the specimens
illustrated by Dimitrov et al. (2022). This is herein considered an intraspecific
variation.

Distribution. Known from Western Anatolia (Dimitrov et al. 2022).

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Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Tegenaria tekke Brignoli, 1978
Figs 18E,.25D—F

Tegenaria tekke Brignoli, 1978b: 516, fig. 98 (9).

Material. TURKIYE: Antalya Prov.: * 3 2 (ZMUU), Kas Dist., Kas-Elmali rd., 916 m,

Pinus brutia and Quercus sp. forest, 20.05.2012 (R.S. Kaya); * 2 2 (ZMUT), same.
Comment. This species was previously known only from its original description.
Distribution. Known only from Antalya Province, southwestern Turkiye.

Discussion

As a result of this study, new taxonomic and faunistic data on the agelenid
spiders of Turkiye, Georgia, and Armenia were provided. Turkiye is one of the
most diverse countries in regards to Agelenidae, with 77 currently known spe-
cies (including the results of the present study). This diversity is indeed higher
compared to several other countries and regions, for example, the entire Cauca-
sus (48 species), Greece (49 species), Bulgaria (44 species), Italy (58 species),
France (41 species), and Spain (41 species) (Nentwig et al. 2024).

In this paper, four new species of Tegenaria were described, including two from
Turkiye and one each from Georgia and Armenia. There are now 39 known species
of this genus in Turkiye (Danigman et al. 2024; present study), and 32 from the Cau-
casus (Otto 2022; Ponomarev et al. 2024). The number of Tegenaria species re-
corded in each Caucasian subregion/country is as follows: Adygea (8), Armenia (1),
Azerbaijan (15), Chechnya (3), Dagestan (7), Georgia (9 or 10), Ingushetia (1), Kab-
ardino-Balkaria (0), Karachay-Cherkessia (3), Krasnodar Krai (13), North Ossetia-Al-
ania (3), South Ossetia (3), and Stavropol Krai (6) (Otto 2022; Ponomarev etal. 2024).

Turkiye has been relatively well studied in terms of its Tegenariini, although
new species and records continue to be discovered regularly. In the present
study, all newly described species from Turkiye were collected from the Tau-
rus Mountain range, a biodiversity hotspot located between the Mediterranean
coastal region and the central Anatolian Plateau (Noroozi et al. 2019). The Tau-
rus Mountains feature high altitudes, diverse valley slopes, depressions, and
rugged karstic landforms, which create a variety of microhabitats and localized
ecological conditions (Atalay et al. 2014). These mountain ranges in Anatolia,
especially those divided by numerous valleys in the south, play a significant
role in speciation and define various biogeographical subregions and provinc-
es. From a zoological perspective, the Anatolian Taurus exhibits a very high de-
gree of endemism and restricted local distributions (Ciplak 2003). This pattern
of distribution underscores the importance of topography and microhabitat
diversity in the evolution and distribution of Tegenaria in Anatolia. Many Ana-
tolian species of Tegenaria are endemics, with most having limited, localized
ranges primarily found in mountainous regions (Kaya et al. 2010). For instance,
the fact that the two closely related species T. bayrami Kaya, Kunt, Marusik &
Ugurtas, 2010 and T. ballarini sp. nov. were both collected sympatrically from
the same habitat further highlights the role of this region in speciation of this
group. The Tegenariini of Georgia and Armenia remain less studied compared
to Turkiye, although the region's diverse habitats and topography suggest that
many undocumented species are still waiting to be discovered.

ZooKeys 1218: 251-286 (2024), DOI: 10.3897/zookeys.1218.135249 282

Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Figure 25. Epigyne of Tegenaria hoeferi sp. nov. (A-C) and T. tekke (D-F). A, D intact, ventral view B, E macerated, ventral
view C, F vulva, dorsal view. Scale bars: 0.2 mm.

Acknowledgments

We thank Dragomir Dimitrov (Sofia, Bulgaria) for providing information on the
variability of the palpal structures in T. percuriosa, and Peter J. Schwendinger
(MHNG) for photographing the holotype of T. hamid. Finally, we thank the
reviewers Mykola Kovblyuk (Crimea, Ukraine) and Martina Pavlek (Zagreb,
Croatia) for their comments and suggestions, from which the manuscript
benefitted greatly.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

ZooKeys 1218: 251-286 (2024), DOI: 10.3897/zookeys.1218.135249 283

Alireza Zamani et al.: New data on the Agelenidae of Turkiye and the Caucasus

Ethical statement

No ethical statement was reported.

Funding

The research of Alireza Zamani was supported by a grant from the Turku University
Foundation (ID 081820).

Author contributions

All authors have contributed equally.

Author ORCIDs

Alireza Zamani © https://orcid.org/0000-0002-8084-9666
Rahsen S. Kaya ® https://orcid.org/0000-0002-3769-9105
Yuri M. Marusik © https://orcid.org/0000-0002-4499-5148

Data availability

All of the data that support the findings of this study are available in the main text.

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